Global negative vegetation feedback to climate warming responses of leaf litter decomposition rates in cold biomes

Whether climate change will turn cold biomes from large long-term carbon sinks into sources is hotly debated because of the great potential for ecosystem-mediated feedbacks to global climate. Critical are the direction, magnitude and generality of climate responses of plant litter decomposition. Here, we present the first quantitative analysis of the major climate-change-related drivers of litter decomposition rates in cold northern biomes worldwide. Leaf litters collected from the predominant species in 33 global change manipulation experiments in circum-arctic-alpine ecosystems were incubated simultaneously in two contrasting arctic life zones. We demonstrate that longer-term, large-scale changes to leaf litter decomposition will be driven primarily by both direct warming effects and concomitant shifts in plant growth form composition, with a much smaller role for changes in litter quality within species. Specifically, the ongoing warming-induced expansion of shrubs with recalcitrant leaf litter across cold biomes would constitute a negative feedback to global warming. Depending on the strength of other (previously reported) positive feedbacks of shrub expansion on soil carbon turnover, this may partly counteract direct warming enhancement of litter decomposition.

中国植物功能型-生物群区的划分

气候在大尺度上决定着植被的分布、结构和组成，植被结构和生理状态的改变可以通过改变植被的反射率、粗糙度以及水分通量进而影响气候，这样形成了气候一植被的相互作用。在植被一气候相互作用的研究中，植物功能型是重要的概念和方法，它可以在详尽描述植被生物物理和生理特征的同时，有效削减植被的复杂性。植物功能型的概念和方法已经在植物群落、生态系统的复杂性和功能、古植被和古气候研究，以及陆面过程模型和动态全球植被模型中得到了广泛的应用。但是针对我国植被-气候的相互作用和区域尺度的全球变化研究，还需要一套特定的植物功能型．生物群区体系。 　　本论文根据我国植被生态学和植被分类的研究背景，结合植被．气候相互作用和区域全球变化研究的需要，提出了一套适宜于中国的植物功能型．生物群区划分方案。首先，根据中国植被和气候特征，筛选并确定了影响植被生物物理和生理属性以及植被分布的6个关键的植物功能特征：然后，根据这6个特征，对植物进行功能型划分，得到了29类植物功能型：再根据我国植被的实际情况和研究需要，选定了其中的18类作为我国的植物功能型。这套功能型包括了7类‘树’功能型，6类‘灌木’功能型和5类‘草’功能型，其中含有4类高寒植物功能型，专门用于描述青藏高原的植被分布，并根据需要设置了2类‘裸地，功能型。 　　根据我国气候一植被分布定量关系的相关研究以及BIOME1和Box体系的研究结果，选定7个环境变量作为限制我国植物功能型分布的关键气候因子：最冷月平均气温、最暖月平均气温、大于50C的有效生长积温、大于OºC的有效生长积温、Priestley-Taylor系数（实际蒸散与潜在蒸散的比值）、降水量、最暖月和最冷月平均气温之差。采用半峰宽法初步确定每个植物功能型的环境限定因子取值范围。并根据这套植物功能型及其环境参数建立了适宜于我国的生物群区体系，从而得到了我国的植物功能型-生物群区体系(the Chinese Plant functional Types and Biomes，CNPB)。 为了验证这套植物功能型-生物群区体系，将BIOME1和中国的植物功能型生物群区体系(CNPB)对中国植被在当前气候条件和未来气候情景下分布的模拟结果进行了比较。结果表明，这套体系可以更有效地模拟中国植被在当前和未来气候条件下的分布，特别是对青藏高原植被描述的详细程度有实质性的提高。

中国西部亚热带常绿阔叶林木本植物小枝功能生态学初步研究

常绿阔叶林以其富饶的生物资源、丰富的生物多样性和巨大的生态与环境效益引起了人们越来越大的重视，它的研究已成为国际植被科学界关注的主题之一。我国分布着世界上面积最大的亚热带常绿阔叶林，在世界植被中具有重要地位，它的分布表现出明显的地带性差异，存在着多样的植物群系及其对应的气候特征。但是在植物功能性状领域，与全球范围其它生物群系相比，常绿阔叶林物种的研究较少，其功能性状间、功能性状与环境间的关系尚不清晰。 本研究以常绿阔叶林木本植物的当年生小枝为对象，试图从小枝水平上的生物量分配格局、叶片大小与数量的权衡关系、小枝茎的构型效应、叶片元素化学计量学，以及小枝大小的成本与效益分析等方面，较为系统地揭示小枝水平上的植物功能性状间及其与气候间的关系。因此，在华西雨屏带内部的不同纬度设置峨眉－青城－雷波－平武的温度梯度进行比较，并对有降水差异的川西南偏湿性（雷波）与偏干性常绿阔叶林（西昌）进行对比研究，同时在不同山体进行不同海拔梯度的比较研究。 本文主要研究结果如下： （1）小枝生物量分配格局叶水平上，叶片重－叶柄重（Y轴vs.X轴,下同）呈斜率小于1的异速生长关系，表明叶柄对叶内部的生物量分配影响显著。小枝水平上，叶和茎的生物量以及它们与小枝总生物量间基本呈等速生长关系，表明大的小枝或大叶物种不一定在叶生物量的分配上占优势。不同生活型间，在小枝或者茎的生物量一定时，常绿物种叶片的生物量比例较落叶物种稍高。与温度和水分较优越（峨眉及其低海拔）的生境相比，在相对低湿（螺髻）与低温（平武）的生境中的植物会减少对叶的投入而增加对支撑部分的投资比例。 （2）小枝叶片大小与数量的权衡无论是不同气候带还是不同生活型以及不同海拔梯度，叶片大小与出叶强度基本都是呈负的等速生长关系，表明了叶片大小－数量在小枝水平上的权衡。在不同气候梯度的对比中，叶片数量（出叶强度）一定时，高温和高水分生境（峨眉）比低温（平武）和低湿（螺髻山）生境中的物种的叶片大小（质量和面积）更大，表明不同生境的比较中，小的叶片可能具有较高的出叶强度和更高的适合度收益。“出叶强度优势”（Leafingintensitypremium）假说可能不适宜解释不同生境物种叶片大小差异。 （3）小枝茎的构型效应虽然茎长和茎径与叶片大小都呈正相关关系，与出叶强度都呈负相关关系，但茎长/茎径比与叶/茎生物量之比呈负相关关系；与叶片的大小呈负相关关系，与出叶强度呈正相关关系。这说明小枝构型能影响小枝叶/茎生物量分配和叶大小－数量的权衡关系。其影响机制可能是小枝内部的顶端优势。另外，茎长/茎径比在低湿和低温等不利生境中的植物中较高，而在降水和温度较适宜环境中较低。 （4）叶片C、N、P化学计量学N含量和P含量，C/N比和比叶重（LMA,leafmassperarea）呈正的等速生长关系，而N和LMA，P和LMA呈负的等速生长关系。在LMA一定时，C/N比随着生境胁迫压力的增加而降低，N、P含量随着生境压力的增加而增加。在P含量一定时，N含量随着生境压力的增加而降低，即N/P比在生境条件较优（峨眉及其低海拔）时较高。常绿和落叶植物叶片的N/P比没有差异，在LMA一定时，常绿植物的N、P含量较高、C/N比较低。总之，植物的C、N、P化学计量学特征受叶片属性如LMA与气候，及其相互作用的影响。 （5）小枝大小的代价与效益分析、TLA与小枝总重总叶面积（TLA,totalleafarea，Y轴，下同）与总叶重（X轴）均呈斜率小于1的异速生长关系，TLA与小枝横切面积呈斜率为1的等速生长关系。表明叶片面积的增加总是小于叶重和小枝总重的增加，随着小枝的增大，它的叶面积支撑效率下降。在热量和降水优越的生境（峨眉及其低海拔）中，相同小枝重或者相同茎横切面积的小枝，其叶面积支撑效率较低湿与低温环境下（螺髻山、平武及高海拔）的高。 总体上，本文初步研究了小枝水平上可能存在的以下三种权衡关系：叶－茎生物量分配权衡；叶片大小－数量的权衡；小枝茎长－茎径的权衡关系，以及气候要素等对这三种权衡关系的影响。在此基础上，我们还讨论了这些权衡关系的可能形成机制，及其与物种生态适应的联系。本研究丰富了生活史对策中关于权衡关系的研究内容，为我国常绿阔叶林功能生态学研究积累了材料。 Evergreen broad-leaved forests are attracting much more attention from vegetation ecologists than ever before because of their abundant nature resource and biological diversity, and also great ecological benefits. China has the largest distribution of subtropical evergreen broad-leaved forests (temperate rainforests) that are typical and representative in the world. The forests span over more than ten degrees in latitude and more than 30 degrees in longitude, providing an ideal place to study plant functional ecology, i.e., the climatic effect on plant functional traits and the relationship between the traits. However, relative to the other biomes, there are few studies addressing functional ecology of the plant species from subtropical evergreen broad-leaved forests. In this study, I focused on the leaf size-twig size spectrum of the woody species of subtropical evergreen broad-leaved forests in southwestern china. I collected data on leaf size and number, twig size in terms of both mass and volume, and stem architecture from five temperate mountains, and then I analyzed the relationships between leaf and stem biomass and between leaf size and number, the effect of stem length/diameter ratio on biomass allocation and on the relationship between leaf size and number, leaf C:N:P stoichiometry, and the twig efficiency of supporting leaf area in relation to twig size. I also addressed the climate effect on the spectrum. The temperature gradient from warm to cool sites was represented by Emei Mountain, Qingchengshan, Leibo, and Pingwu, and the rainfall gradient was assumed to emerge from the comparison between Leibo (High) and Luojishan (Low). In addition, altitudinal effects were analyzed with comparisons between low and high altitudes for each mountains. My main results are as follows. Isometric relationships were found between leaf mass and twig mass and between lamina mass and twig mass, suggesting that the biomass allocation to leaves or laminas was independent of twig mass. Petiole mass disproportionably increase with respect to lamina mass and twig mass, indicating the importance of leaf petioles to the within-twig biomass allocation. In addition, the investigated species tended to have a larger leaf and lamina mass, but a smaller stem mass at a given twig mass at favorable environments including warm and humid sites or at low altitude than unfavorable habitats, which might be due to the large requirements in physical support and transporting safety for the species living at unfavorable conditions. Moreover, the evergreen species invested more in leaves and laminas than the deciduous at given stem or twig biomass within any specified habitats. Negative, isometric scaling relationships between leaf number and size broadly existed in the species regardless of climate, altitude, and life forms, suggesting a leaf size/number trade-off within twigs. Along the climatic gradients, at given leaf number or leafing intensity, the leaves were larger in the favorable environments than the poor habitats. This suggested that the fitness benefit gained by small leaves could be larger than that with high leafing intensity in the stressful sites. I concluded that the “leafing intensity premium” hypothesis was not appropriate to interpreting between-habitat variation in leaf size. Both stem length and diameter were positively correlated to leaf size but negatively correlated to leafing intensity. The ratio of stem length to diameter was negatively correlated to leaf mass fraction, and it was negatively correlated to leaf size but positively correlated to leafing intensity. This suggested that the stem architecture influenced twig biomass allocation and the relationship between leaf size and number. The mechanism underlying the architectural effect might lie in the apical dominance within twig. Moreover, the ratio was greater in unfavorable habitats but smaller in favorable environments. Positive, isometric relationships were found between N and P contents per leaf mass, and between C/N ratio and leaf mass per area (LMA), but N and P contents scaled negatively to LMA. C/N ratio decreased but N and P increased with increasing habitat stress at a given LMA. N content declined with increasing habitat stress at given P content. These indicated that N/P and C/N were higher but LMA was lower in favorable habitats than in the other circumstances. The evergreen and deciduous species were non-heterogeneous in N/P, but the evergreen species have higher N and P contents and lower C/N than the deciduous ones. In general, C:N:P stoichiometry were related to both climatic conditions and other important functional traits like LMA. Total leaf area (TLA) allometricly scaled to leaf mass with a slope shallower than 1, similar to the relationship between TLA and total twig mass (leaf mass plus stem mass), suggesting that TLA failed to keep pace with the increase of leaf mass and twig size. However, TLA scaled isometricly to twig cross-sectional area. Thus, it could be inferred that the twig efficiency of displaying leaf area decreased with increasing twig size. In addition, the efficiency at a given twig size was large in favorable than unfavorable habitats. In general, in this preliminary study, I studied three tradeoff relationships within twigs, i.e., between leaf and stem biomass, between leaf number and size, and between stem length and diameter, as well as the climatic effect on the relationships. I discussed the mechanisms underlying the tradeoff relationships in view of biophysics and eco-physiology of plants. I believe that this study can serve as important materials advancing plant functional ecology of subtropical forest and that it will improve the understanding of life history strategies of plants from this particular biome.